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Zool. Anz. 239 (2000): 147-178 © by Urban & Fischer Verlag ZOOLOGISCHER http://www.urbanfischer.de/journals/zoolanz ANZEIGER The Mandible of Silverfish (Insecta: Zygentoma) and Mayflies (Ephemeroptera): Its Morphology and Phylogenetic Significance1 Arnold H. STANICZEK Eberhard-Karls-Universität Tübingen, Zoologisches Institut, Lehrstuhl für Spezielle Zoologie, Tübingen, Germany Abstract. The mandibles of several mayfly larvae have been investigated in order to reconstruct the groundplan of the mandibular muscles and of the mandibular articulation of Ephemeroptera. The results indicate the presence of three points of mandibular attachment to the cranium in the groundplan of Ephemeroptera. The posterior point of articulation in mayflies corresponds to the posterior (primary) mandibular joint of other Dicondylia. The remaining two points of attachment together form an anterior articulation complex made up of an anterolateral and a postero• medial part. These attachment points of the mayfly mandible are compared with the condition in silverfish. It is shown that in Zygentoma there is a similar anterior articulation complex present. This is interpreted as a groundplan character of Dicondylia. The posteromedial part of this complex in both Zygentoma and Ephemeroptera probably is the homologue of the anterior (secondary) mandibular joint of the remaining Dicondylia. The anterolateral part is secondarily modified in mayflies, where it forms a device to withstand the chewing pressure during adduction of the mandible. The primary mandibular articulation in both mayflies and silverfish is elongated and located dorsally to the secondary one, which results in an oblique axis of mandibular movement. In contrast to this plesiomorphic ar• rangement, the remaining Dicondylia have developed an almost horizontal axis of mandibular movement as well as a tight ball-and-socket type of mandibular articulation. As a consequence of the tightened mandibular articulation, the anterior articulation complex has been simplified in these groups, and a subgenal ridge is developed. The man• dibular musculature of Ephemeroptera and Zygentoma is plesiomorphic in retaining several mandibulo-tentorial muscles as well as two cranial abductor muscles. In all other pterygote lineages the dorsal tentorial muscles are lost, and only a single cranial abductor muscle is present. The mandibular muscles of the silverfish Tricholepidion gertschi show additional plesiomorphies, namely the lack of a mandibulo-hypopharyngeal muscle and the retention of a ligamentous transverse tendon. The groundplan of the dicondyle insect mandible is reconstructed, and the phy• logenetic significance of this character complex is discussed. The mandibles of both Zygentoma and Ephemeroptera do not show any traces of segmentation, thus the assumption of a telognathic insect mandible thereby cannot be supported. The distribution of all investigated mandibular characters support a sistergroup rela• tionship between Ephemeroptera and the remaining pterygote orders, the Metapterygota. Key words. Dicondylia, phylogeny, phylogenetic systematics. 1. INTRODUCTION ent results and conclusions regarding the mandibular articulation. The subimagines and imagines of mayflies do not con• SNODGRASS (1950, 1951) recognized just one mandi• tinue feeding, and their mandibles as well as their other bular joint in the mayfly mandible, whereas other work• mouthparts are highly atrophied in these life stages. ers notified two mandibular articulations (BÖRNER However, the aquatic larvae still bear well-developed 1909; KUKALOVÁ-PECK 1985, 1991; SCHÖNMANN 1981) mandibles. Several authors have investigated the head in different species. Some authors even stated three morphology of mayfly larvae, but came to quite differ• separate articulations (ARENS 1989; BROWN 1961; STRENGER 1953, 1970, 1975, 1977, 1979). 1 This paper is dedicated to Professor Dr. Karl-Ernst Lauter- Similarily, descriptions of silverfish mandibles vary bach (Bielefeld, Germany). considerably. While all authors agree on the presence of 0044-5231/00/239/02-147 $ 12.00/0 148 A. H. STANICZEK an anterior articulation, SNODGRASS (1950, 1952, 1960) The larvae of the New Zealand genus Nesameletus mainly repeatedly referred to the anterior condyle as being lo• feed on diatoms (own observation). cated at the gena, whereas CHAUDONNERET (1950) ob• served an anterior condyle on the laterodorsal angle of • Ameletopsis perscitus Eaton, 1899 (Ameletopsidae) the clypeus. The New Zealand species of the amphinotic distributed In recent publications some authors also claim the Ameletopsidae belongs to one of the very few mayfly fami• presence of a (at least vestigial) segmentation of the lies with larvae with exclusively carnivorous habits. Amele- topsids are known to engulf other water insects (CAMPBELL mandible in both Zygentoma (KRAUS 1998; KUKALOVÁ- 1985). The gut contents of the larger specimens used for this PECK 1998) and Ephemeroptera (KUKALOVÁ-PECK study consisted mainly larvae of Nesameletus and caddisfly 1998). Interpreting the mandible as composed of the larvae. entire telopodite, they revived the dispute between A. perscitus was chosen for this study to observe the impact of MANTON's (1964) idea of a 'whole-limb jaw' in the a totally different nutrition on the anatomy of the mandible. groundplan of Tracheata and the concept of a gnathoba- sic mandible in these groups (LAUTERBACH 1972a). These obvious inconsistencies in both description and 2.2. Additionally used material for interpretation of the mandible in mayflies and silverfish comparisons led to the present study. It is aimed to reconstruct the condition of the mandible in the respective ground- Zygentoma: plans, and to present new arguments for the still contro• Nicoletiidae: Cubacubana spec. versially discussed early phylogenetic branching events Lepismatidae: Lepisma saccharina Linné, 1758 of the Dicondylia. Ctenolepisma spec. Ephemeroptera: 2. MATERIALS AND METHODS Ameletidae: Ameletus inopinatus Eaton, 1887 Metrehtus balcanicus (Ulmer, 1920) 2.1. Specimens examined Ametropodidae: Ametropus fragilis Albarda, 1878 Baetidae: Baetis rhodani (Pictet, 1843) Zygentoma Baetis vernus Curtis, 1834 • Tricholepidion gertschi Wygodzinsky, 1961 Centroptilum luteolum (Müller, 1776) (Lepidothrichidae) Coloburiscidae: Coloburiscus humeralis (Walker, 1853) Ephemerellidae: Serratella ignita (Poda, 1761) This North American relic silverfish is known to possess Torleya major (Klapalek, 1905) some unique plesiomorphic traits not only within Zygentoma Heptageniidae: Ecdyonurus venosus (Fabricius, 1775) (WYGODZINSKY 1961), but also regarding the groundplan of Leptophlebiidae: Habroleptoides confusa Sartori & Jacob, Dicondylia (KRISTENSEN 1998). For instance it is the sole 1986 known recent species of all dicondyle insects that has retained Metretopodidae: Metretopus borealis (Eaton, 1901) the mandibular transverse ligament, otherwise well known Oligoneuriidae: Oligoneuriella rhenana (Imhoff, 1852) from Archaeognaths and entognath insect orders (BOUDRE- Oniscigastridae: Oniscigaster distans Eaton, 1886 AUX 1979a). However, its entire mandibular musculature as Polymitarcyidae: Ephoron virgo (Olivier, 1791) well as its mandibular articulations have not previously been Potamanthidae: Potamanthus luteus (Linne, 1767) subject to a close examination. Rallidentidae: Rallidens mcfarlanei Penniket, 1966 Siphlaenigmatidae: Siphlaenigma janae Penniket, 1962 Ephemeroptera Siphlonuridae: Siphlonurus croaticus Ulmer, 1920 Siphlonurus lacustris (Eaton, 1870) • Oniscigaster wakefieldi McLachlan, 1873 (Oniscigastridae) This species represents one of the two New Zealand represen• tatives of a mayfly family with amphinotic distribution. Its Odonata: imago is considered to resemble the mayfly groundplan in Aeshnidae: Aeshna cyanea Müller, 1764 many aspects (MCLACHLAN 1873; PENNIKET 1962; KLUGE et Calopterygidae: Calopteryx virgo Linne, 1758 al. 1995), so it seemed worthwhile to have also a closer look at its larva. The larva feeds mainly on detritus (MCLEAN 1970). Plecoptera: • Nesameletus sp. (Nesameletidae) Taeniopterygidae: Taeniopteryx spec. Nesameletidae is another mayfly taxon with amphinotic dis• tribution. Its phylogenetic relationship within Ephemeroptera Ensifera: is up to now not satisfactory resolved (KLUGE et al. 1995). Gryllidae: Acheta domesticus Linné, 1767 The Mandible of Silverfish and Mayflies 149 2.3. Methods M16 dorsal suspensor muscle of stomodaeum (M. vertico- pharyngealis) Fixation. The material used was preferably fixed by a mixture M17 lateral dilatator muscle of pharynx (Dilatator pharyn- of 95% ethanol, 35% formaldehyde, and acetic acid (66:33:10). gis lateralis primus) After 24 hours or longer it was transferred to 80% ethanol. Some M18 suspensor muscle of tentorium species (Ametropus, Metretopus) were fixed in ethanol only. M19 antennal muscles (Mm. antennarum) Manual dissection. Larvae were dissected under 80% etha• M20 tentorial adductor muscle of cardo (M. tentoriocardi- nol on a layer of paraffin in a Petri dish. To observe muscles, nalis) they were stained with basic fuchsin. To observe cuticular M21 tentorial adductor muscle of stipes (M. tentoriostipi- structures, specimens were kept in 10% potassium hydroxide talis) under room temperature for several days, until the soft tissues M22 cranial adductor muscle of galeolacinia (M. cranio- dissolved. Then the cuticle was stained with Chlorazol Black. lacinialis) To duplicate the mandible movements, the mandible as well M23 tentorial connective muscle as
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  • Evolution of the Insects

    Evolution of the Insects

    CY501-C11[407-467].qxd 3/2/05 12:56 PM Page 407 quark11 Quark11:Desktop Folder:CY501-Grimaldi:Quark_files: But, for the point of wisdom, I would choose to Know the mind that stirs Between the wings of Bees and building wasps. –George Eliot, The Spanish Gypsy 11HHymenoptera:ymenoptera: Ants, Bees, and Ants,Other Wasps Bees, and The order Hymenoptera comprises one of the four “hyperdi- various times between the Late Permian and Early Triassic. verse” insectO lineages;ther the others – Diptera, Lepidoptera, Wasps and, Thus, unlike some of the basal holometabolan orders, the of course, Coleoptera – are also holometabolous. Among Hymenoptera have a relatively recent origin, first appearing holometabolans, Hymenoptera is perhaps the most difficult in the Late Triassic. Since the Triassic, the Hymenoptera have to place in a phylogenetic framework, excepting the enig- truly come into their own, having radiated extensively in the matic twisted-wings, order Strepsiptera. Hymenoptera are Jurassic, again in the Cretaceous, and again (within certain morphologically isolated among orders of Holometabola, family-level lineages) during the Tertiary. The hymenopteran consisting of a complex mixture of primitive traits and bauplan, in both structure and function, has been tremen- numerous autapomorphies, leaving little evidence to which dously successful. group they are most closely related. Present evidence indi- While the beetles today boast the largest number of cates that the Holometabola can be organized into two major species among all orders, Hymenoptera may eventually rival lineages: the Coleoptera ϩ Neuropterida and the Panorpida. or even surpass the diversity of coleopterans (Kristensen, It is to the Panorpida that the Hymenoptera appear to be 1999a; Grissell, 1999).